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Investigating the origin of hazelnut

(Corylus avellana L.) cultivars using
chloroplast microsatellite
ARTICLE in GENETIC RESOURCES AND CROP EVOLUTION SEPTEMBER 2009
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Genet Resour Crop Evol (2009) 56:851859

DOI 10.1007/s10722-009-9406-6

RESEARCH ARTICLE

Investigating the origin of hazelnut (Corylus avellana L.)

cultivars using chloroplast microsatellites
Paolo Boccacci Roberto Botta

Received: 14 July 2008 / Accepted: 12 January 2009 / Published online: 31 January 2009

Springer Science+Business Media B.V. 2009

Abstract The place and time of European hazel

(Corylus avellana L.) domestication is not clear,
although it was already cultivated by the Romans. In
this study, 75 accessions from Spain, Italy, Turkey,
and Iran were analysed using 13 chloroplast microsatellite to investigate the origin and diffusion of
hazelnut cultivars. Four loci were polymorphic and
identified a total of four different chlorotypes. Their
distribution was not uniform in each geographical
group. The most frequent chlorotype A was present in
all groups. An increase in chlorotype number and
diversity from Spain eastward to Italy, Turkey, and
Iran was observed. Results suggest that some spread
of cultivars occurred from East to West and that
hazelnut cultivation was not introduced from the
eastern Mediterranean basin into Spain and southern
Italy by Greeks or Arabs. Moreover, the results
suggest considerable exchange of germplasm
between Italy and Spain, probably by the Romans.
Hazelnut appears to have been domesticated independently in three areas: the Mediterranean, Turkey,
and Iran.

P. Boccacci (&)
R. Botta

Dipartimento di Colture Arboree, Universita` degli Studi di
Torino, Via Leonardo da Vinci 44, 10095 Grugliasco,
Torino, Italy
e-mail: paolo.boccacci@unito.it

Keywords Chlorotype
Corylus avellana

cpSSR
Cultivar diffusion
Cultivation

Domestication
Filbert

Introduction
In Europe, two different Corylus species are present:
the European hazel, C. avellana L., that has a wide
distribution, and the Turkish hazel, C. colurna L.,
restricted to the Balkans, Romania, and northern
Turkey (Thompson et al. 1996). The geographical
distribution of the European hazel extends from the
Mediterranean coast of North Africa northward to the
British Islands and the Scandinavian peninsula, and
eastward to the Ural Mountains of Russia, the
Caucasus Mountains, Iran, and Lebanon (Kasapligil
1972). It is the source of important cultivars in Europe
and Turkey, which were selected over many centuries
from local wild populations (Trotter 1921; Tasias
Valls 1975; Thompson et al. 1996). Turkey has long
been the leading producer and exporter of hazelnuts,
accounting for about 71% of world production. Italy is
second with over 13%, the United States third with
4.1%, and Spain fourth with 2.8%. Azerbaijan, Iran,
Georgia, China, France, and Greece are other producers (FAOstat 2008). About 90% of the world crop
is used as kernels in the food industry, while the
remaining 10% is sold in-shell for fresh consumption.
The present-day distribution of C. avellana L. was
established about 7,000 years B.P., as result of a

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852

postglacial recolonization process starting approximately 11,000 years earlier (Huntley and Birks 1983).
During the most recent glaciation hazel was restricted
to glacial refugia in southern Europe, but low levels of
pollen were deposited in central Europe during the full
glacial period (Bennett et al. 1991). Huntley and Birks
(1983) suggested that southern Italy and the area
around the Bay of Biscay (southwestern France) were
the most important refugia. Between 10,000 and
9,000 years B.P. the amount of Corylus pollen found
across Europe sharply increased (http://www.
pierroton.inra.fr/Cytofor/Maps/index.html). The most
plausible explanation for this increase is a very rapid
spread of hazel (1,500 m/year) from these refugia
(Huntley and Birks 1983; Birks 1989). Recent analysis
of chloroplast DNA variation (Palme and Vendramin
2002) indicates a rapid expansion from one large
refugium in the Biscay area or from several scattered
refugia in western Europe into most of Europe, except
southern-central Italy and the Balkans, and then a local
expansion in the latter two areas.
Nut dispersal during the postglacial recolonization
was caused by animals (small mammals and birds)
and human migration. A prevalent opinion is that
Mesolithic tribes (10,0006,000 years B.P.) could
have aided, intentionally or more likely accidentally,
the spread of hazel. However, there is no evidence
that they attempted to propagate it (Tallantire 2002).
In several Mesolithic archaeological findings,
abundant nutshell fragments have been recorded,
indicating that hazelnuts were cracked for consumption or some kind of nut-processing (Bakels 1991;
Kubiak-Martens 1999). Most likely the expansion of
the hazelnut distribution was due to chance spread
during the preparation of hazelnut meals by
migratory Mesolithic people (Kuster 2000). During
the spread of agriculture in Europe, C. avellana L.
was one of the species domesticated and cultivated
(Forni 1990; Zohary and Hopf 2001). Archaeologists
have repeatedly retrieved nuts, kernels, and shell
remains from many Neolithic, Bronze Age, Classical,
and Medieval sites all over Europe (Bakels 1991;
Russel-White 1995; Arobba et al. 2003; Cleary 2003;
Pena-Chocarro et al. 2005; Sostaric et al. 2006;
Schmidl et al. 2007). Nevertheless, where and when
the domestication of C. avellana L. was started is not
yet clear; although it was cultivated by the Romans
(Trotter 1921; White 1970; Vaughan and Geissler
1997).

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Genet Resour Crop Evol (2009) 56:851859

In recent years, the availability of nuclear microsatellite or simple sequence repeat (nSSR) markers in
C. avellana L. (Bassil et al. 2005a, b; Boccacci et al.
2005) provide new possibilities to investigate the
breeding history of hazelnut cultivars. nSSR loci have
been used to fingerprint and to identify mistakes in
hazelnut accessions from several germplasm repositories (Botta et al. 2005; Gokirmak et al. 2008). These
studies investigated genetic relationships among
cultivars grown in important production areas,
identified accessions with identical fingerprints, and
suggested the parentage for several cultivars
(Boccacci et al. 2006, 2008; Ghanbari et al. 2005;
Gokirmak et al. 2008). In particular, Boccacci et al.
(2006) and Gokirmak et al. (2008) concluded that
cultivars from Italy and Spain are genetically related.
On the contrary, Turkish cultivars constitute a
separate germplasm group, indicating little gene flow
between the eastern and western areas of the
Mediterranean basin.
The chloroplast genome has a lower evolutionary
rate than the nuclear genome. Although its inheritance is paternal in conifers (Vendramin and
Ziegenhagen 1997), inheritance is maternal in angiosperms (Dumolin et al. 1995; Arroyo-Garca et al.
2002). Thus in angiosperms the chloroplast genome
can only be disseminated by seeds or cuttings.
Chloroplast SSR (cpSSR) markers have been developed in recent years (Provan et al. 2001) and have
been used to investigate the origin and diffusion of
several fruit tree species, such as grape (ArroyoGarca et al. 2002; Imazio et al. 2005; Arroyo-Garca
et al. 2006) and olive (Breton et al. 2006).
In the present work, a total of 75 genotypes from four
different geographical regions were analysed using
cpSSR markers to investigate the history of hazelnut
cultivation and diffusion. Moreover, the usefulness of
cpSSRs as molecular markers to study genetic relationships among hazelnut cultivars was evaluated.

Materials and methods

Plant material and DNA extraction
Seventy-five hazelnut accessions were sampled in
field collections assembled by the Dipartimento di
Colture Arboree of Torino (Italy); Cooperativa San
Colombano (Genova, Italy); Istituto di Frutti-

Genet Resour Crop Evol (2009) 56:851859

viticoltura of Piacenza (Italy); Consiglio per la

Ricerca e Sperimentazione in Agricoltura (CRA) of
Roma and Caserta (Italy); National Clonal Germplasm Repository (NCGR) of Corvallis (Oregon,
USA), Institut de Recerca i Tecnologia Agroalimenta`ries (IRTA) of Constant (Tarragona, Spain),
and Seed and Plant Improvement Institute (SPII) of
Kamal-Abad (Iran). Trueness-to-type of the accessions was previously verified by Boccacci et al.
(2005, 2008) and Ghanbari et al. (2005) using nSSR
markers. The cultivars were assigned to four groups
on the basis of their country of origin (Table 1).
Cultivars with a direct parental relationship were
excluded, as were synonyms. (i) Group Spain is
composed by 33 cultivars grown in the province of
Tarragona (Catalonia, northeastern Spain), except
Casina, which is from Asturias (northern coast of
Spain); (ii) Group Italy includes 22 cultivars grown in
different Italian regions: Piedmont and Liguria
(northwestern Italy), Latium (central Italy), and
Campania and Sicily (southern Italy); (iii) Group
Turkey is represented by ten cultivars of Turkish
origin, seven grown in the Black Sea coastal provinces of northern Turkey, and three cultivated in
Greece (Extra Ghiaghli, Sivri Ghiaghli, and
Tombul Ghiaghli); (iv) Group Iran is composed of
10 accessions from Iran.
Total genomic DNA was extracted from 0.2 g of
leaves or immature catkins using the modified
procedure described by Thomas et al. (1993) in a
TrisEDTANaCl buffer containing 0.25 M NaCl,
0.2 M Tris pH 7.6, 2.5% PVP 40,000, 0.05 M
Na2EDTA, and 0.1% b-mercaptoethanol. After purification, the DNA was suspended in 50 ll Tris
EDTA buffer.
cpSSR analysis and data elaboration
A total of thirteen cpSSR loci were analysed. Eight
loci (ccmp1, ccmp2, ccmp3, ccmp4, ccmp5, ccmp6,
ccmp7, ccmp10) were described by Weising and
Gardner (1999), and five loci (cmcs1, cmcs2, cmcs4,
cmcs11, cmcs13) were described by Sebastiani et al.
(2004). The primer pairs were those designed by
these authors for Nicotiana tabacum L. and Castanea sativa Mill., respectively. PCR amplification was
carried out using a reaction mixture (20 ll) consisting
of 50 ng DNA template, 0.5 lM of each primer,
200 lM dNTPs, 2 mM MgCl2, 2 ll 109 NH4 buffer

853

[160 mM (NH4)2SO4, 670 mM TrisHCl (pH 8.8 at

25C), 0.1% Tween-20], and 0.5 U BioTaq DNA
polymerase (Bioline, London, UK). All cpSSR
amplifications were performed using the following
temperature profiles: 3 min of denaturation at 95C,
then 28 cycles of 30 s of denaturation at 95C, 45 s of
annealing at 55C, and 90 s of extension at 72C; and
a final 10 min extension step at 72C. Amplified
fragments were loaded onto a polyacrylamide gel and
run on a semi-automated ABI-PRISM 377 sequencer
(Applied Biosystems, Foster City, Calif., USA).
Results of the run were processed with Genescan
software and allele sizes (in base pairs, bp) were
estimated using a GeneScan-350 ROX size standard
(Applied Biosystems).
Numbers of chlorotypes (N) were directly estimated for each cultivar group. An unbiased estimate
of the
was calculated as:
 chlorotype diversity


H n=n  1 1  Rg2i , where gi is the frequency of the i-th chlorotype and n is the number
of individuals analysed in each cultivar groups (Nei
1987). A chlorotype median network was constructed
using the program Network v. 4.5 (Bandelt et al.
1999).

Results
cpSSR polymorphism and chlorotype definition
In the 75 hazelnut genotypes, nine of 13 cpSSR loci
were monomorphic. Low levels of variation were
detected with the other four loci (Table 2). Locus
ccmp2 showed three different size variants, whereas
two variants were found at loci ccmp3, ccmp4, and
ccmp10. The allele variations differed by increments
of 1 bp due to variation in the number of A or T
residues within mononucleotide repeats. The same set
of cpSSR loci was also polymorphic in 26 natural
hazel populations (Palme and Vendramin 2002).
Thus, ccmp2, ccmp3, ccmp4, and ccmp10 are useful
loci for studying genetic variability in C. avellana L.
Considering the allele variants at the four loci, four
different chlorotypes were detected (Table 3) and
their relationships were analysed under a network
model (Fig. 1). The chlorotype network indicated the
minimum number of evolutionary events separating
each chlorotype. Chlorotype A was the most frequent
in the 75 cultivars. Chlorotypes B and D were

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854

Genet Resour Crop Evol (2009) 56:851859

Table 1 List of accessions, germplasm collections, and chlorotypes observed in 75 hazelnut cultivars
Collectiona

Chlorotype

Cultivar

Ametllenca

IRTA

Pinyolenc

IRTA

Apegalos
Artellet

IRTA
IRTA

A
A

Planeta
Punxenc

IRTA
IRTA

A
A

Casina

DCA

Queixal de llop

IRTA

Castanyera (syn. Barcelona)

DCA

Queixal de ruc

IRTA

Closca molla
Culpla`

DCA

IRTA

DCA

Ratllada
Ratol`

IRTA

Gironell (syn. Grossal)

DCA

Ribet

IRTA

Gironenc (syn. Vermellet)

NCGR

Ros

IRTA

Grifoll

IRTA

Rosset

IRTA

Lluenta

IRTA

Sant Jaume

IRTA

Martorella

IRTA

Sant Joan

IRTA

Morell

DCA

Sant Pere

IRTA

Cultivar

Collectiona

Chlorotype

Group Spain (33 cultivars)

Negret

IRTA

Segorbe

IRTA

Negret capellut
Negret garrof`

IRTA

Trenet

IRTA

IRTA

Vimbod`

IRTA

Pauetet

DCA

Group Italy (22 cultivars)

Camponica

DCA

Nocchione

DCA

Catainetto

Coop

Noscello

Coop

Del Rosso

Coop

Nociara

DCA

DellOrto

Coop

Riccia di Talanico

CRA

Ghirara

DCA

San Giovanni

DCA

Gianchetta

IFP

Tonda bianca

DCA

Iannusa Racinante

DCA

Tonda di Giffoni

DCA

Menoia

IFP

Tonda Gentile delle Langhe

DCA

Mortarella

DCA

Tonda Gentile Romana

CRA

Napoletana

DCA

Tonda rossa

CRA

Napoletanedda

CRA

Trietta

IFP

Badem

IRTA

Palaz

DCA

Extra Ghiaghli

IRTA

Sivri

IRTA

Imperial de Trebizonde (syn. Karidaty)

Incekara

DCA
IRTA

B
B

Sivri Ghiaghli
Tombul

NCGR
IRTA

B
A

Kalinkara

NCGR

Tombul Ghiaghli

NCGR

Asle Gharebag

SPII

Pashmineh

SPII

Dobooseh

SPII

Rasmi

SPII

Jorow Gharebag

SPII

Shastak-2

SPII

Mish-pestan

SPII

Shirvani

SPII

Group Turkey (10 cultivars)

Group Iran (10 cultivars)

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Genet Resour Crop Evol (2009) 56:851859

855

Table 1 continued
Cultivar

Collectiona

Chlorotype

Cultivar

Collectiona

Chlorotype

Nakhoni Rood

SPII

Tabari Rood

SPII

DCA, Dipartimento di Colture Arboree of Torino (Italy); IRTA, Istitut de Recerca i Tecnologia Agroalimenta`ries of Reus
(Tarragona, Spain); Coop, Cooperativa San Colombano (Genova, Italy); IFP, Istituto di Frutticoltura of Piacenza (Italy); CRA, Centro
Ricerche e sperimentazione in Agricoltura of Roma and Caserta (Italy); NCGR, National Clonal Germplasm Repository of Corvallis
(Oregon, USA); SPII, Seed and Plant Improvement Institute (Karaj, Iran)

Table 2 Chlorotype genotypes at 13 cpSSR loci

Chlorotype

ccmp1

ccmp2

ccmp3

ccmp4

ccmp5

ccmp6

ccmp7

ccmp10

cmcs1

cmcs2

cmcs4

cmcs11

cmcs13

130

214

119

117

108

98

154

108

103

134

108

224

118

130

214

118

117

108

98

154

108

103

134

108

224

118

C
D

130
130

215
216

118
119

117
116

108
108

98
98

154
154

108
107

103
103

134
134

108
108

224
224

118
118

Only four loci (ccmp2, ccmp3, ccmp4, and ccmp10) were polymorphic and identified four chlorotypes

Table 3 Chlorotype
distribution in each
geographical group (with
chlorotype frequency in
parentheses)

n = number of samples;
N = number of
chlorotypes;
H = chlorotype diversity

Group Spain

Group Italy

Group Turkey

Group Iran

Total

33

22

10

10

75

Chlorotype A

33 (1.000)

20 (0.909)

4 (0.400)

1 (0.100)

58 (0.773)

Chlorotype B

6 (0.600)

1 (0.100)

7 (0.094)

Chlorotype C

8 (0.800)

8 (0.108)

Chlorotype D

2 (0.091)

2 (0.027)

0.000

0.173

0.533

0.378

0.391

associated with chlorotype A, while chlorotype C was

related only to chlorotype B. The intermediate
relationship of chlorotype A to the others suggests
that it is likely the ancestral chlorotype of hazelnut
cultivars.
Chlorotype variation and distribution
in the geographical groups
The chlorotype distribution was not uniform in the
four geographical groups (Table 3). Chlorotype A
was present in all groups with a decreasing frequency
from Spain eastward to Italy, Turkey, and Iran
(Table 3). It was observed in all 33 Spanish and 20
of 22 Italian accessions. Tonda bianca and Tonda
rossa, both of which are cultivated in Avellino
province (Campania, southern Italy), showed the
rarest chlorotype D (Table 1). Chlorotypes B and C
were mainly present in the groups Turkey and Iran,

respectively, and were absent in Spanish and Italian

cultivars (Table 3).
In order to evaluate the chlorotype distribution in
each analysed group, a chlorotype diversity index (H)
was calculated (Table 3). H values were zero in the
Spanish group, and low in the Italian group. On the
contrary, high H values were observed in groups
Turkey (H = 0.533) and Iran (H = 0.378), although
a relatively fewer accessions from these groups were
analysed. These data showed that H increases from
West to East in the Mediterranean basin, unlike what
was observed in grapevine by Imazio et al. (2005).

Discussion
The postglacial migration of C. avellana L. was
hypothesized on the basis of pollen records (Huntley
and Birks 1983; Birks 1989) and a recent analysis of

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856

Fig. 1 Chlorotype median network representing all chlorotypes (A, B, C, and D) identified in hazelnut. Circle areas are
proportional to chlorotype frequencies obtained in all 75
samples analysed

chloroplast DNA variation in wild hazelnuts (Palme

and Vendramin 2002). The most likely scenario is an
expansion from southwestern France into most of
Europe, except for Italy and the Balkans where
expansions were local. Archaeobotanical remains
indicate that hazel kernels have been an important
constituent in the human diet since prehistoric times.
Kernels are tasty and have a high energy value (600
680 kcal per 100 g fresh weight) due to their high fat
and protein content. In addition, hazelnuts are easy to
store and transport. Mesolithic tribes could have
aided, intentionally or more likely accidentally, the
spread of hazel. According to Tallantire (2002), they
did not attempt to propagate it.
The place and time of hazel domestication is not
clear. The common opinion is based on a statement of
Pliny the Elder (2379 A.D.) in the work Naturalis Historia that the hazelnut came from Asia
Minor and Pontus (north coast of Turkey). On the
basis of this assertion, the accepted general idea is
that hazelnut cultivation was brought to Italy by the
Greeks and that the specific epithet avellana derives
from Abellina in western Asia, allegedly the present

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Genet Resour Crop Evol (2009) 56:851859

valley of Damascus (Trotter 1921; Rosengarten

1984). Contacts and trade between eastern and
western Mediterranean basin as long ago as the
second half of the XVI century B.C. are documented
through abundant archaeological findings dating back
to the Mycenaean civilization (15001100 B.C.). The
colonization of the Italian peninsula by the Greeks
(Magna Graecia) may have contributed to transfer of
the cultivation techniques and cultivars of grape,
olive, and other species to the local populations and
eventually the Romans (Forni 1990).
The cpSSR results of our analysis show an
increase in chlorotype number and diversity from
West to East. In addition, a decrease in frequency of
the most common chlorotype A from the groups
Spain and Italy to Turkey and Iran was observed
(Table 3). These results suggest little gene flow from
East to West, and that Spanish and Italian accessions
have a common genetic base. The nSSR analyses of
Boccacci et al. (2006) and Gokirmak et al. (2008)
support this hypothesis, as both studies indicated a
close genetic relationship between Spanish and
Italian cultivars. On the contrary, Turkish cultivars
were assigned to a separate cluster, indicating little
gene flow between the eastern and western areas of
the Mediterranean basin. Thus, both cpSSR and nSSR
data indicate that hazelnut cultivation and cultivars
were not introduced from the eastern Mediterranean
basin into Spain and southern Italy by Greeks or by
Arabs (Trotter 1921; Alberghina 2002). According to
Alfonso (1886) and other Sicilian authors, hazelnut
cultivation was introduced to Sicily by Arabs. Yet
Arabs only dominated the island after the second half
of the IX century A.D., whereas hazelnut was already
cultivated in Campania (southern Italy) centuries
earlier by the Romans (Trotter 1921).
Evidence of the importance of hazelnut cultivation
during the Roman civilization was reported in several
Latin classics (Trotter 1921). In Catos (234149
A.D.) De re rustica the hazelnut was mentioned as
nux avellana, and the author recommended cultivation on farms of nuces, calvas, avellanas,
praenestinas et graecas. In Columellas (I century
A.D.) De re rustica and Plinys (2379 A.D.)
Naturalis Historia this species was reported in the
list of the plants that grown on farms. In the poetic
texts (Eclogues and Georgics) of Virgil (7019 B.C.),
hazelnut was mentioned in a mountainous context,
often in association with pastoral activities, as well as

Genet Resour Crop Evol (2009) 56:851859

in large-scale agricultural production. In Palladiuss

(IVV century A.D.) Opus agriculturae a multiplication technique of hazelnut was briefly described.
Hazelnut was also mentioned in the alimentary or
medicinal Latin literature. Dioskurides (I century
A.D.) recommended the use of kernels as a cure for
the common cold and persistent coughs (Rosengarten, 1984). Apicius (1437 A.D.) in the manuscript
De re coquinaria reported a recipe for the preparation
of a dessert similar to a nougat using hazelnuts and
honey (Trotter 1921). The geographical region which
most Latin authors mention in relation to the
cultivation of hazelnut was Campania. The latifundium, the entrepreneurial management of agricultural
estates using groups of slaves, started after the II
century B.C. in Latium and Campania regions, and
then became widespread under Roman rule (Sirago
1995). On the other hand, the Latin world adopted the
terms abellanae and abellinae to indicate the hazelnut tree and its fruit, respectively, from which is
derived the specific epithet avellana. These expressions derived from Abella and Abellinum, the actual
towns of Avella and Avellino in Campania, respectively, where hazel was widely cultivated during
Roman times. Servius (IIIIV century A.D.), in a
comment to the Georgics poem by Virgil, reported:
sane coryli proprie dicuntur, nam abellanae ad
Abellano (Abellino in other documents) Campaniae
oppido, ubi abundant, nominatea sunt (Trotter
1921). Other evidence of the importance of hazelnut
production in this region during the Roman period are
the archaeological remains and the pollen data. The
presence of carbonized hazelnuts and the representation of hazel trees and nuts in wall paintings found in
the excavations at Pompeii and Herculaneum (Trotter
1921; Jashemski 1970), indicate that hazelnuts were
consumed and commercialized also in areas relatively distant from the traditional cultivation centers.
Pollen maps indicate that in 2500 B.P. the Corylus
pollen percentage was low in southern Italy (05%),
but it increased to 510% around 2000 B.P. and to
1015% around 1500 B.P. (http://www.pierroton.
inra.fr/Cytofor/Maps/index.html), most likely in
relation to an increase in human activity.
The historical documents and archaeological findings indicate that hazelnut cultivation and
consumption were significant during the Roman
period. This suggests that the use of hazelnut was
very likely spread throughout the empire by the

857

Romans. The presence of chlorotype A in all

analysed groups (Table 3) indicates a common origin
of the accessions; the intermediate relationship of
chlorotype A to the others suggests that it could be an
ancestral chlorotype (Fig. 1). Thus, it can be hypothesized that Italian hazelnut germplasm was spread by
the Romans to the Iberian peninsula (150100 B.C.)
and to a lesser extent to Asia Minor (133 B.C.) during
their expansion. Moreover, cpSSR and nSSR
(Boccacci et al. 2006) results suggest that these
exchanges were stronger and frequent between Italy
and Spain. In the province of Tarragona (Catalonia,
northeastern Spain), where 88% of the Spanish
hazelnut area is located, Roman influence was strong.
The Romans improved existing cities, such as
Tarragona (Tarraco), and superimposed the latifundium on the local landholding system. This supports
the hypothesis that the Romans may have introduced
the idea of systematically cultivating and using
hazelnut in their dominions. Human migrations and
trade between the central and western Mediterranean
basin were also frequent after the Roman civilization,
because southern Italy was dominated by the Spanish
from the XIV to the XVIII century. The prevalence of
chlorotype A in Spanish and Italian accessions
suggests that southern Italy, most likely the Campania
region, was an important centre of origin and diffusion
of hazelnut cultivars. The archaeological findings,
historical document, and pollen data already discussed
support this hypothesis. In addition, the absence
of chlorotypes B and C in the groups Spain and
Italy suggests that hazelnut varieties were domesticated separately in Turkey and Iran, with perhaps
limited germplasm exchange with Italy and Spain.
In conclusion, our results show the usefulness of
chloroplast markers to study relationships among
cultivars. The chlorotype distribution allowed us to
hypothesize a possible geographical origin and diffusion of hazelnut cultivars. Moreover, the
integration of genetic data with historical, archaeological, and palynological information indicates that
the history of C. avellana cultivation in Europe is
similar to that of Castanea sativa Mill. In fact, as
observed in chestnut by Conedera et al. (2004), our
results indicate a very limited influence of the Greek
hazelnut cultivation techniques on the Latin and
subsequent Roman civilization, unlike the situation in
grape and olive. Thus, it is probable that systematic
cultivation and use of hazelnut was spread by the

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858

Romans, probably from the Campania region. We

investigated few cultivars from Turkey and Iran, but
found considerable chlorotype diversity. Our data
point toward separate domestications in these
regions, and limited germplasm exchange with the
western to eastern Mediterranean basin. Further
analyses of wild populations and cultivars from other
geographical areas, using chloroplast and nuclear
markers, would improve our understanding of the
origin and spread of hazelnut cultivars.
Acknowledgements Authors are grateful to: Dr. Nahla V.
Bassil (USDA-ARS National Clonal Germplasm Repository of
Corvallis, Ore.), Dr. Roberto De Salvador and Dr. Pasquale
Piccirillo (Consiglio per la Ricerca e sperimentazione in
Agricoltura, Italy), Dr. Silvia Dellepiane (Cooperativa San
Colombano, Genova, Italy), Dr. Alireza Ghanbari (Department
of Horticultural Science, Shahed University, Iran), Dr. Merce
Rovira (Istitut de Recerca i Tecnologia Agroalimenta`ries,
Tarragona, Spain), and Dr. Virginia Ughini (Istituto di Fruttiviticoltura, Universita` Cattolica S.C., Piacenza, Italy) for
providing samples of leaves or catkins. Authors thank prof.
Shawn A. Mehlenbacher (Oregon State University, Corvallis,
Ore.) for reviewing the manuscript. The research was funded
by Regione Piemonte Administration (CIPE, 2003) and MIUR
(ex 60%, 2006).

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